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Osteological Evidence for Paraphyly of the Avian Order Caprimulgiformes (Nightjars and Allies) Gerald Mayr

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Osteological Evidence for Paraphyly of the Avian Order Caprimulgiformes (Nightjars and Allies) Gerald Mayr J. Ornithol. 143, 82±97 (2002) ã Deutsche Ornithologen-Gesellschaft/Blackwell Wissenschaftsverlag, Berlin ISSN 0021-8375 Osteological evidence for paraphyly of the avian order Caprimulgiformes (nightjars and allies) Gerald Mayr Forschungsinstitut Senckenberg, Sektion fuÈr Ornithologie, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; Email: [email protected] Summary The first phylogenetic analysis of osteological characters in the Caprimulgiformes (night- jars and allies) provides strong evidence that this order is paraphyletic. Well supported is monophyly of a clade comprising Caprimulgidae, Nyctibiidae, Aegothelidae and the tradi- tional Apodiformes (swifts and hummingbirds). Within this clade Nyctibiidae (potoos) and Caprimulgidae (nightjars) on the one hand, and Aegothelidae (owlet-nightjars) and the traditional Apodiformes on the other are sister taxa. Whereas monophyly of Nyctibiidae and Caprimulgidae is in accordance with most previous studies, monophyly of the so far poorlystudied Aegothelidae and the traditional Apodiformes has not been suggested by previous authors. The affinities of the Steatornithidae (oilbirds) and Podargidae (frog- mouths) cannot be conclusivelyresolved at present. Keywords: Phylogeny, Caprimulgiformes, Apodiformes, Aegothelidae, osteology. Zusammenfassung Osteologische Hinweise auf Paraphylie der Vogelordnung Caprimulgiformes (SchwalmvoÈgel) Die erste phylogenetische Analyse der Caprimulgiformes (SchwalmvoÈgel) aufgrund osteologischer Merkmale liefert deutliche Hinweise auf eine Paraphylie dieser Ordnung. Gut gestuÈtzt ist die Monophylie eines Taxons, das Caprimulgidae, Nyctibiidae, Aegotheli- dae und die traditionelle Ordnung Apodiformes (Segler und Kolibris) beinhaltet. Inner- halb dieser Gruppe sind Nyctibiidae (TagschlaÈfer) und Caprimulgidae (Ziegenmelker) auf der einen Seite, sowie Aegothelidae (HoÈhlenschwalme) und die traditionellen Apodifor- mes auf der anderen Seite monophyletisch. WaÈhrend Monophylie von Nyctibiidae und Ca- primulgidae im Einklang mit den meisten bisherigen Untersuchungen steht, wurde eine Monophylie der bisher kaum untersuchten Aegothelidae und der traditionellen Apodifor- mes noch nicht vorgeschlagen. Die groûsystematische Stellung der Steatornithidae (Fett- schwalme) und Podargidae (Eulenschwalme) kann zur Zeit nicht uÈberzeugend begruÈndet werden. Introduction 19th and early20th century,well before the concepts of modern phylogenetic systematics Virtuallyall of the currentlyrecognized avian were introduced (Hennig 1950), and some re- orders were established byanatomists in the main verypoorlydiagnosed. An especiallyill- U.S. Copyright Clearance Center Code Statement: 0021-8375/2002/14301±0082 $ 15.00/0 G. Mayr ´ Paraphyly of Caprimulgiformes 83 defined order is the Caprimulgiformes which closelyrelated (see Sibley& Ahlquist 1990: traditionallyincludes five recent families: pp. 414±418 for a detailed review of the his- the holarctic Caprimulgidae (nightjars), the toryof classification). Cracraft (1981: 700) neotropic Nyctibiidae (potoos) and Steatorni- noted some morphological characters in order thidae (oilbirds), and the Australasian Aego- to support this traditional classification. How- thelidae (owlet-nightjars) and Podargidae ever, the characters he listed as synapomor- (frogmouths). All are crepuscular or nocturnal phies of Aegothelidae and Podargidae are ei- birds, and most feed on flying insects (the ther absent in stem-group Podargidae Steatornithidae are frugivorous). Caprimulgi- (ªdouble-notched sternum with long posterior form birds have a superficiallysimilar external processesº, see Mayr 1999, 2001), incorrect appearance which includes a wide beak with a (ªanterior iliac blades extremelywell devel- verylarge gape, and a soft plumage with cryp- opedº ± in the podargid genus Batrachostomus tic colouration. In their osteologyand internal the alae praeacetabulares ilii are verynarrow), anatomy, however, the members of this order or insufficientlydefined (ªpeculiar egg-white are remarkablydifferent. Monophylyof the protein patternº). In a recent paper, Cracraft Caprimulgiformes has never been convinc- (2001: Fig. 3) considered the systematic posi- inglyestablished with derived morphological tion of Aegothelidae and Podargidae to be un- characters, and the characters usuallylisted to resolved. diagnose this order (e. g. Stresemann 1927±34, Wetmore (1960) thought the Steatornithidae Sibley& Ahlquist 1990) are either not present to be the sister taxon of all other caprimulgi- in all taxa, plesiomorphic (e. g., anisodactyl form birds, which is reflected bytheir classifi- feet, holorhinal nares, presence of aftershaft), cation into a separate suborder. Cracraft (1981, or present in a wide range of other birds (e. g., 2001) assumed that this familyis the sister the pelvic muscle formula, arrangement of the taxon of Nyctibiidae and Caprimulgidae but, flexor tendons, nude oil gland). Despite this again, the characters he listed in order to sup- obvious lack of autapomorphic characters, the port monophyly of Steatornithidae, Nyctibii- traditional order Caprimulgiformes was con- dae, and Caprimulgidae are either not present sidered monophyletic by all recent and most in all three taxa (contraryto his statement, the earlier authors. Among the few exceptions are caudal margin of the sternum of the Nyctibii- Huxley(1867) who separated the Podargidae dae does not bear a single lateral process but from Nyctibiidae, Caprimulgidae, and Aego- exhibits two pairs of incisions) or plesiomor- thelidae, and Sharpe (1900) who separated phic (presence of processus basipterygoidei, Steatornithidae, Podargidae, and Aegothelidae see below). Curiously, Cracraft (1988: 354) from Nyctibiidae and Caprimulgidae in his lin- stated that ªthe steatornithids are seemingly ear classification. [. .] primitive relative to the other four fami- The phylogenetic relationships between the liesº. caprimulgiform families also are poorlyre- Recent molecular phylogenetic studies do solved. Most authors based their studies on not support either monophyly of Caprimulgi- comparisons with Steatornithidae, Podargidae, dae and Nyctibiidae, or monophyly of Podargi- and Caprimulgidae, whereas published ac- dae and Aegothelidae. The DNA-DNA hybrid- counts of the anatomyof Nyctibiidaeand Ae- ization studies of Sibley& Ahlquist (1990) gothelidae are exceedinglyrare. Although only suggested monophyly of the taxon (Caprimul- Cracraft (1981) presented a formal phyloge- gidae + (Nyctibiidae + Steatornithidae)), with netic hypothesis based on morphological char- the Aegothelidae as the most basal taxon of re- acters, generallyNyctibiidaeand Caprimulgi- cent Caprimulgiformes. A phylogenetic analy- dae on the one hand, and Aegothelidae and sis using the mitochondrial cytochrome b gene Podargidae on the other were considered to be byMariaux & Braun (1996) showed the taxon 84 Journal fuÈr Ornithologie 143, 2002 (Nyctibiidae + (Caprimulgidae + Aegotheli- teros (2000) did confirm monophyly of Capri- dae)) to be monophyletic with the Steatornithi- mulgiformes and Apodiformes in most of the dae as the sister taxon of all other Caprimul- different trees that were presented (in the anal- giformes. Both studies supported an early yses of Bleiweiss et al. 1994, Mindell et al. branching of the Podargidae (which branch 1997, and Espinosa de los Monteros 2000 only just after the Aegothelidae respectivelyStea- the Caprimulgidae were included). tornithidae in the phylogenies of Sibley & Ahl- Further strong evidence for a close relation- quist and Mariaux & Braun). An analysis by ship between the apodiform and (at least some) Johansson et al. (2001) using nuclear DNA se- caprimulgiform birds comes from the well quences did not corroborate monophyly of known osteologyof the Aegialornithidae, an Steatornithidae, Podargidae, Nyctibiidae, and Eocene familyof basal swifts. The Aegialorni- Caprimulgidae (the Aegothelidae were not in- thidae (which are considered to be apodiform cluded in the analysis). However, the proposed byvirtuallyall recent authors, e. g. Olson phylogenetic positions of these taxa received 1985, Peters 1985, Karhu 1988, Mourer-Chau- onlyweak statistical support and are thus not vire 1988a) not onlycloselyresemble the re- discussed in the following. cent Caprimulgidae and Aegothelidae in their Concerning their higher systematic position osteology(see Olson 1985 and below), but are most authors considered the Caprimulgiformes also verysimilar to the roughlycontemporane- to be most closelyrelated either to owls (Stri- ous extinct caprimulgiform familyArchaeo- giformes) or to swifts and hummingbirds trogonidae (Mourer-Chauvire 1980, 1995). On (Apodiformes) (see Sibley& Ahlquist 1990). the other hand, there are no derived similarities However, morphological similarities between between earlyTertiaryowls and contempo- caprimulgiform birds and owls have mainly raneous caprimulgiform birds, nor derived been found if comparisons were made with Po- morphological characters which support dargidae and Steatornithidae (e. g., FuÈrbringer monophyly of owls and the traditional order 1888, Verheyen 1956), whereas the other fami- Caprimulgiformes. lies share derived characters with the Apodi- I attempted to analyze the exact phyloge- formes. Although the DNA-DNA hybridiza- netic relationships between apodiform and tion studies of Sibley& Ahlquist (1990) and caprimulgiform birds. The proposed relation- Bleiweiss et al. (1994) also resulted in mono- ships are shown in a cladogram, all nodes of phyly of the taxon
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